Petr Jansta

The circumscription of the family Ormyridae (Hymenoptera: Chalcidoidea) is revised after phylogenetic analysis based on ultra‐conserved elements (UCEs) and comparative morphological assessment of the chalcid ‘Gall Clade’. Six genera are treated in the family, including two new genera, Halleriaphagus van Noort and Burks, gen. nov ., and Ouma Mitroiu, gen. nov. One genus, Eubeckerella Narendran, is re‐assigned to the family, and Ormyrulus Bouček is synonymised with Ormyrus Westwood, syn. nov ., resulting in the new combination Ormyrus gibbus (Bouček), comb. nov. The six genera are classified in three subfamilies, two of which are newly described, Asparagobiinae van Noort, Burks, Mitroiu and Rasplus, subfam. nov., and Hemadinae van Noort, Burks, Mitroiu and Rasplus, subfam. nov. Halleriaphagus is established for the newly described type species Halleriaphagus phagolucida van Noort and Burks, sp. nov ., and Ouma is erected for O. daleskeyae Mitroiu, sp. nov. , and O. emazantsi Mitroiu, sp. nov. Asparagobius is revised with description of Asparagobius bouceki van Noort, sp. nov. , and Asparagobius copelandi Rasplus and van Noort, sp. nov. Asparagobius and Halleriaphagus are classified in Asparagobiinae, Hemadas in Hemadinae and Eubeckerella , Ormyrus and Ouma in Ormyrinae. The molecular support defining the ormyrid clade is corroborated by the proposed morphological synapomorphy of a foliaceous prepectus overlying the tegula base. Identification keys to the genera of Ormyridae and to the species of Asparagobius and Ouma are provided. Online Lucid identification keys and images of all the species treated herein are available at: http://www.waspweb.org . Zoobank Registration: LSID urn:lsid:zoobank.org:pub:8811695B-EE57-4C18-A6B6-E63D267E2373 .

Chalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra‐conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first‐instar larvae. The phylogeny suggests a transition from smaller soft‐bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the “Angiosperm Terrestrial Revolution”. Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3–180.5 Ma) and a crown age of 162.2 Ma (153.9–169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history.